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Ganong's Review of Medical Physiology, 23rd Edition

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316 SECTION IV Endocrine & Reproductive <strong>Physiology</strong><br />

and glucagon excess makes diabetes worse. Excess pancreatic<br />

production <strong>of</strong> somatostatin causes hyperglycemia and other<br />

manifestations <strong>of</strong> diabetes.<br />

ISLET CELL STRUCTURE<br />

The islets <strong>of</strong> Langerhans (Figure 21–1) are ovoid, 76- × 175μm<br />

collections <strong>of</strong> cells. The islets are scattered throughout the<br />

pancreas, although they are more plentiful in the tail than in<br />

the body and head. β-islets make up about 2% <strong>of</strong> the volume<br />

<strong>of</strong> the gland, whereas the exocrine portion <strong>of</strong> the pancreas (see<br />

Chapter 26) makes up 80%, and ducts and blood vessels make<br />

up the remainder. Humans have 1 to 2 million islets. Each has<br />

a copious blood supply; blood from the islets, like that from<br />

the gastrointestinal tract (but unlike that from any other endocrine<br />

organs) drains into the hepatic portal vein.<br />

The cells in the islets can be divided into types on the basis<br />

<strong>of</strong> their staining properties and morphology. Humans have at<br />

least four distinct cell types: A, B, D, and F cells. A, B, and D<br />

cells are also called α, β, and δ cells. However, this leads to<br />

confusion in view <strong>of</strong> the use <strong>of</strong> Greek letters to refer to other<br />

structures in the body, particularly adrenergic receptors (see<br />

Chapter 7). The A cells secrete glucagon, the B cells secrete<br />

insulin, the D cells secrete somatostatin, and the F cells<br />

secrete pancreatic polypeptide. The B cells, which are the<br />

most common and account for 60–75% <strong>of</strong> the cells in the<br />

islets, are generally located in the center <strong>of</strong> each islet. They<br />

tend to be surrounded by the A cells, which make up 20% <strong>of</strong><br />

the total, and the less common D and F cells. The islets in the<br />

tail, the body, and the anterior and superior part <strong>of</strong> the head<br />

<strong>of</strong> the human pancreas have many A cells and few if any F<br />

cells in the outer rim, whereas in rats and probably in<br />

humans, the islets in the posterior part <strong>of</strong> the head <strong>of</strong> the pancreas<br />

have a relatively large number <strong>of</strong> F cells and few A cells.<br />

FIGURE 21–1 Islet <strong>of</strong> Langerhans in the rat pancreas. Darkly<br />

stained cells are B cells. Surrounding pancreatic acinar tissue is lightcolored<br />

(× 400). (Courtesy <strong>of</strong> LL Bennett.)<br />

A variety <strong>of</strong> other hormones also have important roles in<br />

the regulation <strong>of</strong> carbohydrate metabolism.<br />

The A-cell-rich (glucagon-rich) islets arise embryologically<br />

from the dorsal pancreatic bud, and the F-cell-rich (pancreatic<br />

polypeptide-rich) islets arise from the ventral pancreatic<br />

bud. These buds arise separately from the duodenum.<br />

The B cell granules are packets <strong>of</strong> insulin in the cell cytoplasm.<br />

The shape <strong>of</strong> the packets varies from species to species;<br />

in humans, some are round whereas others are rectangular<br />

(Figure 21–2). In the B cells, the insulin molecule forms polymers<br />

and also complexes with zinc. The differences in the<br />

shape <strong>of</strong> the packets are probably due to differences in the size<br />

<strong>of</strong> polymers or zinc aggregates <strong>of</strong> insulin. The A granules,<br />

which contain glucagon, are relatively uniform from species<br />

to species (Figure 21–3). The D cells also contain large numbers<br />

<strong>of</strong> relatively homogeneous granules.<br />

STRUCTURE, BIOSYNTHESIS, &<br />

SECRETION OF INSULIN<br />

STRUCTURE & SPECIES SPECIFICITY<br />

Insulin is a polypeptide containing two chains <strong>of</strong> amino acids<br />

linked by disulfide bridges (Table 21–1). Minor differences<br />

occur in the amino acid composition <strong>of</strong> the molecule<br />

from species to species. The differences are generally not<br />

sufficient to affect the biologic activity <strong>of</strong> a particular insulin<br />

FIGURE 21–2 Electron micrograph <strong>of</strong> two adjoining B cells<br />

in a human pancreatic islet. The B granules are the crystals in the<br />

membrane-lined vesicles. They vary in shape from rhombic to round<br />

(× 26,000). (Courtesy <strong>of</strong> A Like. Reproduced, with permission, from Fawcett DW:<br />

Bloom and Fawcett, A Textbook <strong>of</strong> Histology, 11th ed. Saunders, 1986.)

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